The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands

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Zookeys 578: | 15—160 (2016)

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The tiger beetles (Coleoptera, Carabidae,
Cicindelinae) of Israel and adjacent lands

Andrey V. Matalin'?, Vladimir I. Chikatunov?

| Education-Scientific Centre Ecology & Biodiversity, Moscow State Pedagogical University, Moscow 129164,
Russia 2 Department of Biology, Russian National Research Medical University named after N.I. Pirogov,
Moscow 117997, Russia 3 Department of Zoology, Tel-Aviv University, Tel Aviv 69978, Israel

Corresponding author: Andrey V. Matalin (a_matalin@tochka.ru; andrei-matalin@yandex.ru)

Academic editor: A. Casale | Received 2 December 2015 | Accepted 12 February 2016 | Published 8 April 2016
http://zoobank.org/[ALA7FC2B-0E 1D-4BC8-8AE4-30CC9478DF7B

Citation: Matalin AV, Chikatunov VI (2016) The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and
adjacent lands. ZooKeys 578: 115-160. doi: 10.3897/zookeys.578.7383

Abstract

Based on field studies, museums collections and literature sources, the current knowledge of the tiger
beetle fauna of Israel and adjacent lands is presented. In Israel eight species occur, one of them with two
subspecies, while in the Sinai Peninsula nine species of tiger beetles are now known. In the combined re-
gions seven genera from two tribes were found. The Rift Valley with six cicindelids species is the most spe-
cious region of Israel. Cylindera contorta valdenbergi and Cicindela javeti azari have localized distributions
and should be considered regional endemics. A similarity analysis of the tiger beetles faunas of different
regions of Israel and the Sinai Peninsula reveal two clusters of species. The first includes the Great Rift
Valley and most parts of the Sinai Peninsula, and the second incorporates most regions of Israel together
with Central Sinai Foothills. Five distinct adult phenological groups of tiger beetles can be distinguished
in these two clusters: active all-year (three species), spring-fall (five species), summer (two species), spring-
summer (one species) and spring (one species). The likely origins of the tiger beetle fauna of this area are
presented. An annotated list and illustrated identification key of the Cicindelinae of Israel and adjacent

lands are provided.

Keywords
Carabidae, tiger beetles, Cicindelinae, Israel, Lebanon, Jordan, Syria, Egypt, Sinai, Levant, Mediterra-

nean, fauna, endemic, near-endemic, catalogue, key, distribution, phenology, faunogenesis

Copyright A.V. Matalin, V.l. Chikatunov. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

116 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

Introduction

The first data about tiger beetles of Palestine were published in the first third of the
XXth century. In 1913 Sahlberg described from Wadi El Nawaime (modern Wadi
en Nuweima) Cicindela littoralis aulicoides. 1n 1934 Mandl recorded for the Palestine
two subspecies of Cicindela littoralis: C. 1. winkleri and C. Ll. aulicoides. The first species
list of Palestinian Coleoptera including five species of tiger beetles was published by
Bodenheimer in 1937. Around the same time, the first information about cicindelids
of the Sinai Peninsula appeared and Cicindela aulica (Horn, 1931), Cicindela litto-
ralis aulicoides (Mandl, 1934) and Megacephala euphratica (Schatzmayr, 1936) were
recorded. Unfortunately, detailed locality data and collecting dates for specimens of
these species were often incomplete.

A second wave of tiger beetles studies in the Levant was completed in the last third
of XXth century. Alfieri (1976) published the catalogue of Egyptian Coleoptera with
information about 11 species of tiger beetles, six of which were recorded for the Sinai
Peninsula. The first data about Cicindelinae of Israel were published by Valdenberg
(1983, 1985) and Nussbaum (1987). It should be noted that these papers also con-
tained information about tiger beetles of the Sinai Peninsula. In all eight species were
recorded from Israel and seven species for the Sinai. Unfortunately, in the paper by
Nussbaum (1987) data about localities for several species given in the text and on the
maps do not coincide.

Since the beginning of 2000 interest in the Cicindelinae of the Middle East has
increased significantly (El-Moursy et al. 2001; Franzen 2001, 2007; Finkel et al. 2002;
Wiesner 2002, 2005; Abdel-Dayem et al. 2003; Rittner 2003; Abdel-Dayem 2004,
2012; Chikatunov et al. 2006; Avgin and Ozdikmen 2007; Franzen 2007; Avgin and
Wiesner 2009; Ptashkovsky 2009; Deuve 2011, 2012; Abdel-Dayem and Kippenhan
2013; Jaskuta and Rewicz 2014). These studies revealed the presence of several species
of tiger beetle previously unknown from the area. For example, Habrodera nilotica
(Dejean, 1825), Hypaetha singularis (Chaudoir, 1876) and Cephalota littorea (Forskal,
1775) were recorded for the first time in Israel (Chikatunov et al. 2006). However, in
the next publications these species were not included (Ptashkovsky 2009).

During the last decade, new information about the distribution of tiger beetles in
different parts of the Levant has accumulated, and we include these new records here.

Material and methods

Specimens and data for this report come from the following museums and private col-
lections:

TAU Tel Aviv University (Israel);
ZMUM Zoological Museum of Moscow State University (Moscow, Russia);
MPU —_ Moscow State Pedagogical University (Moscow, Russia);

The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands LF

SIZ I.I. Schmalhausen Institute of Zoology, National Academy of Sciences of
Ukraine (Kiev, Ukraine);
cJW collection of Jiirgen Wiesner (Wolfsburg, Germany);

clOv collection of Igor’ Ovsyannikov (Moscow, Russia).

The nomenclature of elytral pattern follows Acciavatti and Pearson (1989); the no-
menclature of male internal sac follows Matalin (1998); the chorology follows Vigna
Taglianti et al. (1999) with some additions; the regions of Israel and the Sinai Penin-
sula (Egypt) follow Nussbaum (1987). The similarity of the faunas of tiger beetles was
calculated using complete linkage procedure (squared Euclidean distances).

The species included here that are not yet recorded from Israel are marked in the
catalogue and in the key with a symbol (0).

Results and discussion

Catalogue of the tiger beetles of Israel and adjacent lands

Family Carabidae Latreille, 1802
Subfamily Cicindelinae Latreille, 1802
Tribe Cicindelini Latreille, 1802
Subtribe Cicindelina Latreille, 1802
Genus Calomera Motschulsky, 1862

Calomera aulica aulica (Dejean, 1831)

General distribution. Europe - Greece; Asia - Lebanon, Israel, Jordan, Syria, Egypt
(Sinai), Saudi Arabia, Arab Emirates, Oman, Yemen, Bahrain, Iran, Iraq, Pakistan;
Africa: Cape Verde Islands, Senegal, Guinea Bissau, Mauritania, Morocco, Tunisia,
Algeria, Libya, Sudan, Chad, Egypt, Somalia, Eritrea, Djibouti.

References. IsraeL — Bodenheimer 1937: 108 (as Cicindela); Valdenberg 1983:
43, 46 (as Cicindela), 1985: 37 (as Cicindela); Cassola 1985: 56 (as Lophyridia); Nuss-
baum 1987: 9-10 (as Cicindela); Wiesner 1992: 151 (as Lophyridia); Puchkov and
Matalin 2003: 99; Rittner 2003 (as Lophyridia); Ptashkovsky 2009: 8-9 (as Lophyra);
EeypT (S1nal) — Horn 1931: 162 (as Cicindela); Alfieri 1976: 1 (as Cicindela); Cassola
1984: 56 (as Lophyridia); Nussbaum 1987: 9-10 (as Cicindela); Wiesner 1992: 151
(as Lophyridia); Werner 2000: 98 (as Lophyridia); El-Moursy et al. 2001: 66 (as Cicin-
dela); Abdel-Dayem et al. 2003: 205 (as Lophyridia); Puchkov and Matalin 2003: 103;
Abdel-Dayem 2004: 74 (as Lophyridia).

Distribution (Figs 1, 2). ISRAEL (INCLUDING STATE OF PALESTINE), Jordan Valley:
Zor Deir Shaman, 15.11.2005, I. Zonstein - 12; Gesher, 17.VU1.1939, H. Bytinski-
Salz - 14 (both TAU); Kinneret zone (after Nussbaum 1987); Dead Sea Area: Ne‘ ot
HaKikkar, 7.V.1980, leg. A. Valdenberg - 553 799; 16.VII.1999, 13.VHI.1999,

118 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

ir) ):
Pivigiel
we

SF ay
* ey,
——
io

Figure |. Distribution of Calomera aulica aulica in Israel, Palestine and border areas of Jordan (open circles
- records before year 1949, half-solid circles - records between years 1950-1999, solid circles - records after
year 2000; map source - Eric Gaba Wikimedia Commons user: Sting and Wikimedia Commons user: Nor-
dNordWest, URL - https://upload.wikimedia.org/wikipedia/commons/7/7c/Israel_relief_location_map.jpg)

HS)

the tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands

Figure 2. Distribution of Calomera aulica aulica (red circles) and Calomera littoralis aulicoides (blue
thombs) in Sinai Peninsula, Egypt (open symbols - records before year 1949, half-solid symbols - between
years 1950-1999, solid symbols - records after year 2000; URL map source - https://upload.wikimedia.

org/wikipedia/commons/5/59/Sinai_relief_location_map.svg).

120 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

11.1X.1999, and 12.XII. 1999, light trap BL, leg. I. Yarom & V. Kravchenko - 2¢'3)
72.2; Sedom, 15.VIII.1957, leg. J. Wahrman - 1¢ 299 (all TAU); “xn Gedi, 19-
29.V.1989, leg. G. Miiler - 12; Qalya, 28.VIII.1986, 28.6.1987, leg. Y. Nussbaum
- 20 (both cJW); “not Qane (after Nussbaum 1987); Arava Valley: Be‘er Ora,
3.1V.1997, leg. V. Chikatunov - 354 19; En Tddan, 15.V11.1999, leg. I. Yarom &
V. Kravchenko - 1¢' 19 (all TAU).

Jorpan, Ma’Daba: Callirhoe, 7.V1.1942, leg. H. Bytinski-Salz - 29. (TAU).

Ecyrt (Srnat), Northern Sinai: Sabkhat al Bardawil, 25.V111.1967, leg. I. Mar-
galit - 39 9; 24.VII1.1979, leg. A. Valdenberg - 1¢ 299 (TAU); Ismailia (after Alfieri
1976); Zaranik Protectorate (after El-Moursy et al. 2001; Abdel-Dayem et al. 2003;
Abdel-Dayem 2004); Sinai Mountains: 20 km NE of Dahab, saline land, 4.VIII.2008,
leg. A. Sokolov - 4¢55292 (MPU); Southwestern Sinai: Suez — 1¢ (ZMUM); Na-
beq, 17.VIII.1971, leg. J. Kugler -13 229; 8.V.1980, leg. A. Valdenberg - 44.4
6929; Ras al Tantur, 5.VII.1957, leg. Ch. Lewinsohn - 2A UD. TNA TOF A leg.
M. Kaplan -19 (all TAU); 15 km W Offra, Golf v. Elat, Strafe von Tiran, 3.I1V.1981,
leg. G. Gerdes - 14 (JW); Wadi Gharandal, 20.V.1969, leg. Tsabar - 13 (TAU);
Abu Zenima, Wadi Tayebeh (both after Alfieri 1976); E/ Tor (after Alfieri 1976; Abdel-
Dayem et al. 2003; Abdel-Dayem 2004); Ras Muhammad (after Nussbaum 1987).

Calomera littoralis aulicoides (J. Sahlberg, 1913)

General distribution. Asia - Turkey, Israel, Jordan, Syria, Egypt (Sinai), Saudi Arabia,
Iran, Iraq; Africa - Egypt.

References. IsraEL — Sahlberg 1913: 3 (as Cicindela); Mandl 1934: 244-245 (as
Cicindela lunulata nemoralis aulicoides), 1982: 93-94 (as Lophyridia aulicoides); Val-
denberg 1983: 44, 47 (as Cicindela), 1985: 36 (as Cicindela); Nussbaum 1987: 11-12
(as Cicindela); Wiesner 1992: 149 (as Lophyridia); Puchkov and Matalin 2003: 100;
Chikatunov et al. 2006: 293; Ecypr (Stnat) - Mandl 1934: 244-245 (as Cicindela
lunulata nemoralis aulicoides), 1982: 94 (as Lophyridia aulicoides); Alfieri 1976: 2 (as
Cicindela lunulata aulicoides); Wiesner 1992: 149 (as Lophyridia); Abdel-Dayem et al.
2003: 207 (as Lophyridia); Puchkov and Matalin 2003: 103; Abdel-Dayem 2004: 74
(as Lophyridia).

Distribution (Figs 2-3). IsRaAEL (INCLUDING STATE OF PALESTINE), Golan
Heights: Hammat Gader, 2.X.2002, leg. V. Kravchenko & V. Chikatunov - 192
(TAU); Lower Galilee: Teverya, 16.VI.1981, leg. A. Valdenberg - 19; Kinneret,
16.V1.1981, leg. A. Valdenberg - 345 329; Jordan Valley: Zor Deir Shaman,
Yarden bank, 32°02'30’'N, 35°30'E, 15.11.2005, leg. L. Friedman & I. Zonstein
- 3535 19 (TAU); Allenby bridge (after Mandl 1982); Tomer, Ma’oz-Hayyim (both
after Nussbaum 1987); Dead Sea Area: Yeriho, Jordan, Palestine, 24.1V.27 — 19;
Enot Zugim, 13.111.1993, leg. V. Chikatunov - 203 19, 9.V1.1997, leg. L. Fried-
man - 14, 1.11.1994, and 13.111.1994, leg. V. Chikatunov - 3¢¢ 299; Ne‘ot
Hakakrar AIA 999: - bo. VU N9995 13. Vile 999 eand e199 9% les. I. Yarom

the tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 124

tt

maT |
A

~~
wit =

Figure 3. Distribution of two subspecies of Calomera littoralis in Israel, Palestine and border areas of
Jordan (red circles — C. 1. aulicoides, blue circles — C. 1 winkleri, open circles - records before year 1949,
half-solid circles — records between years 1950-1999, solid circles — records after year 2000; map source
- Eric Gaba Wikimedia Commons user: Sting and Wikimedia Commons user: NordNordWest, URL -
https://upload.wikimedia.org/wikipedia/commons/7/7c/Israel_relief_location_map.jpg).

122 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

& V. Kravchenko - 173.4119. 9; Sedom, 19.VIII.1957, J. Wahrman - 243; Qalya,
9.VI.1981, leg. A. Valdenberg - 43.3 62 (all TAU); “xn Gedi, 1-13.V.1980, 19-
29.V.1989, leg. G. Miiller - 329; Newe Zohar, 24.V1.1987, leg. Y. Nussbaum -
13 (both cJW); Wadi El Nawaime [Wadi en Nu’eima] (after Sahlberg 1913); Enot
Qane (after Nussbaum 1987); Arava Valley: ‘Ex Zin, 30°53.60'N, 35°09.17'E, light
trap BL, 12.X.1999, leg. I. Yarom & V. Kravchenko - 14; Hazeva, field school,
30°46.70'N, 35°14.25’'E, light trap BL, 20.11.1999, 21.V.1999, leg. I. Yarom &
V. Kravchenko - 14 399; ‘En ‘Iddan, 20.V1.1995, leg. I. Yarom & A. Freidberg -
23 19; 15.VH.1999, leg. I. Yarom & V. Kravchenko —3¢'¢ 499; Nahal Shezaf,
18.V.1999, 8.VI.1999, light trap, leg. I. Yarom & V. Kravchenko - 14 192; Nahal
Negarot, 10.11.1999, leg. I. Yarom & V. Kravchenko - 19 (all TAU).

Jorpan, Al Balqa’: Al Maghtas, 12.11.1942, leg. H. Bytinski-Salz - 1¢299(TAU);
Ma’Daba: Suwayma, Dead Sea, 5.IV.2000, leg. G. & I. Zappi - 1419 (MPU).

Eeyrt (Sinal), Northern Sinai: /smailia (after Alfieri 1976); Southwestern Sinai:
El Tor (after Alfieri 1976; Abdel-Dayem et al. 2003; Abdel-Dayem 2004).

Calomera littoralis winkleri (Mandl, 1934)

General distribution. Europe - Greece, Armenia, Azerbaijan; Asia - Cyprus, Turkey,
Lebanon, Israel, Jordan, Syria, Iran, Iraq, Afghanistan, Turkmenistan.

References. IsraeL — Mandl 1934: 240, 243, 245 (as Cicindela); Naviaux 1983:
82 (as Lophyridia), Valdenberg 1983: 44, 46 (as Cicindela), 1985: 36 (as Cicindela);
Nussbaum 1987: 11, 13 (as Cicindela); Wiesner 1992: 151 (as Lophyridia); Puchkov
and Matalin 2003: 101; Rittner 2003 (as Cicindela); Chikatunov et al. 2006: 293;
Ptashkovsky 2009: 8-9 (as Cicindela).

Distribution (Fig. 3). IsraEL, Upper Galilee: Huda, 23.V1.1952, leg. J. Wahrman
-5353 42 9; 8.11.1976, leg. M. Kaplan - 14; Sasa, 18.111.1951, leg. J. Wahrman - 19
(all TAU); Northern Coastal Plain: ‘Akko, 7.VIII.1980, leg. A. Valdenberg - 303
422; Maagan Mikha‘el, 17-18.111.1979, 27.V1.1979, 26.]I1.1980, 24.1V.1980,
24.V1.1980, leg. A. Valdenberg - 1743 289 9; 4.V1.1983, leg. E. Sney-Dor - 263
12; Nahariyya, 19.V1.1942, leg. H. Bytinski-Salz - 1¢ 299; Dor, Horvat Tantura,
sea-shore, 13.1X.1949, leg. J. Wahrman - 19; Zikhron Ya aqov, 29.V1.1998, leg. A.
Traub - 329 (all TAU); Central Coastal Plain: Hadera, 28.111.2008, leg. G. Wizen
-14 19; Bat Yam, 13.V11.1945, leg. H. Bytinski-Salz - 1¢ 399; Hofit, 21.1X.1994,
leg. F. Kaplan & A. Freidberg - 19; Mishmeret, 3.VUI.1983, leg. A. Freidberg - 243
3292; Qesarya, 11.V11.1979, and 10.VIII.1979, leg. A. Valdenberg - 35'd 49.9; Tel
Aviv, 20.V1.1982, leg. A. Valdenberg - 1¢ 29 9; 12.IV.2003, leg. V. Kravchenko &
V. Chikatunov - 44'¢ 29 9; Rosh Ha Ayin, 16.1V.1993, leg. A. Freidberg & F. Kaplan
- 14 19; Rehovot, 5.V.1942, leg. H. Bytinski-Salz - 354 229 (all TAU); South-
ern Coastal Plain: Nitzanim, 13.VII.1981, leg. A. Valdenberg - 33 ¢ 229 (TAU);
Judean Hills: Jerusalem - 1¢ (after Mandl 1934: 40, Fig. 65).

The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 123

Genus Cephalota Dokhtouroff, 1883
Cephalota (Taenidia) zarudniana vartianorum (Mandl, 1967)

General distribution. Asia - Israel, Syria, Iran, Iraq.

References. IsraEL — Naviaux 1983: 78; Valdenberg 1983: 43 (as Cicindela jarud-
niana vartinorum), 45 (as Cicindela jorudniana), 1985: 37 (as Cicindela jarudniana);
Nussbaum 1987: 9, 13 (as Cicindela jarudniana); Wiesner 1992: 177; Puchkov and
Matalin 2003: 103; Chikatunov et al. 2006: 293 (as Cephalota deserticola); Ptashko-
vsky 2009: 8-9 (as Cephalota deserticola).

Distribution (Fig. 4). IsRaEL (INCLUDING STATE OF PALESTINE), Dead Sea Area:
Yeriho, 24.1V.1927, leg. O. Theodor - 14; Enot Zugim, 1.11.1994, and 13.11.1994,
leg. V. Chikatunov - 15 229; Qalya, 6.V.1980, leg. A. Valdenberg - 25d 399;
Ne‘ ot HaKikkar, 7.V.1980, leg. A. Valdenberg - 44.4; 28.1V.1984, leg. E. Shney-Dor
- 299; 19.IV.1999, leg. I. Yarom & V. Kravchenko - 141 (all TAU), 7.V.1980, leg.
R. Naviaux - 12; V.1990, leg. Orbach - 1419 (both cJW).

Comments. References to Cephalota zarudniana vartianorum (Mandl, 1967)
as Cephalota deserticola (Faldermann, 1836) (Chikatunov et al. 2006; Ptashkovsky
2009) were based on two mis-identified males from Qalya by K. Mandl. According to
Franzen and Wiesner (1998) C. deserticola is distributed in the western part of Central
Asia, as well as in Iran, Azerbaijan, Armenia and north-eastern Turkey.

(0) Cephalota (Taenidia) littorea littorea (Forskal, 1775)

General distribution. Asia - Egypt (Sinai), Saudi Arabia; Africa - Egypt, Sudan, Eritrea.

References. Ecypt (S1nal) - Forskal 1775: 77 (as Cicindela); Alfieri 1976: 2 (as
Cicindela); Valdenberg 1983: 44, 46 (as Cicindela), 1985: 37 (as Cicindela); Nussbaum
1987: 9, 15 (as Cicindela); Gebert 1991: 176, 187; Wiesner 1992: 175; Werner 2000:
147; Abdel-Dayem et al. 2003: 199; Puchkov and Matalin 2003: 103; Rittner 2003;
Abdel-Dayem 2004: 72.

Distribution (Fig. 5). Ecypr (Sina), Sinai Mountains: Dahab, 9.V.1980, leg.
A. Valdenberg - 1¢ 19; Ras-Burka, 5.1X.1976, leg. A. Freidberg - 13 (all TAU);
Sun Pool (after Nussbaum 1987); Southwestern Sinai: Suez (after Forskal 1775; Ge-
bert 1991); Nabeq, 8.V.1980, 17.VIII.1978, 31.V.1980, leg. A. Valdenberg - 2505
24292; 29.V.1981, leg. A. Freidberg - 234 (all TAU); E/ Tor (after Alfieri 1976;
Abdel-Dayem et al. 2003; Abdel-Dayem 2004); Ras Muhammad, 16.VIII.1978, leg.
A. Valdenberg - 244 19; (after Nussbaum 1987; Gebert 1991).

Comments. The specimen of Cephalota littorea littorea (Forskal, 1775) with label
“Jerusalem” from Zoologisches Museum der Humboldt-Universitat (Berlin) is misla-
belled (see Gebert 1991). All subsequent records of this species from Israel (Wiesner
1992; Puchkov and Matalin 2003) are in error.

124 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

Figure 4. Distribution of Cephalota zarudniana vartianorum (orange triangles), Cicindela javeti azari
(red circles) and Cylindera contorta valdenbergi (blue rhombs) in Israel, Palestine and border areas of Leba-
non (open symbols - records before year 1949, half-solid symbols — records between years 1950-1999;
map source - Eric Gaba Wikimedia Commons user: Sting and Wikimedia Commons user: NordNor-
dWest, URL - https://upload.wikimedia.org/wikipedia/commons/7/7c/Israel_relief_location_map.jpg).

the tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 125

Figure 5. Distribution of Cephalota tibialis tibialis (red circles), Cephalota littorea littorea (blue rhombs)
and Habrodera nilotica nilotica (green triangles) in Sinai Peninsula, Egypt (open symbols - records before
year 1949, half-solid symbols — records between years 1950-1999; URL map source - https://upload.

wikimedia.org/wikipedia/commons/5/59/Sinai_relief_location_map.svg).

126 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

In some publications (Abdel-Dayem et al. 2003; Abdel-Dayem 2004) Cephalota
circumdata (Dejean, 1822) was recorded from the Sinai Peninsula (El Tor). How-
ever, the nominotypical subspecies of C. circumdata occurs along the Aegean, Mar-
mora, Black and Mediterranean Sea costs in the Greece, Bulgaria, western Turkey,
and, probably Rumania (Franzen 1996; Cassola 1999; Gebert 1999); C. c. cappadocica
Franzen, 1996 and C. c. hattusae Franzen, 1996 live along banks of the salt lakes in the
central Turkey (Franzen 1996; Cassola 1999; Gebert 1999); C. c. leonschaeferi Cassola,
1970 occupies the Mediterranean sea cost in southern France (including Corsica) and
north-western Italia (Gebert 1999); while C. c. imperialis Klug, 1834 records in the
Italia (Sardinia and Sicilia), south-eastern Spain (including Balearic Islands), Tunisia
and Algeria, but not in the Libya and Egypt (Gebert 1999). Most likely, the aberrant

specimen of C. /ittorea was incorrectly identified as C. circumdata.

(0) Cephalota (Taenidia) tibialis tibialis (Dejean, 1882)

General distribution. Asia - Egypt (Sinai); Africa - Egypt.

References. Ecypt (Srna) — Valdenberg 1983: 42, 47 (as Cicindela); Nussbaum
1987: 7, 12 (as Cicindela), 1985: 37 (as Cicindela); Gebert 1991: 179, 187; Wiesner
1992: 175; El-Moursy et al. 2001: 66 (as Cicindela littorea); Abdel-Dayem et al. 2003:
200; Puchkov and Matalin 2003: 103; Abdel-Dayem 2004: 72, 2012: 198.

Distribution (Fig. 5). Ecypr (Srnat), Northern Sinai: Yamit, 21.V1.1978,
14.VII.1981, leg. A. Valdenberg — 322 (TAU); Sabkhat al Bardawil, 7.V1.1977,
26.VII.1978, 31.VIII.1978, 7.VI.1980, leg. A. Valdenberg - 4394722 (TAU); Arish
(after Abdel-Dayem et al. 2003); Zaranik Protectorate (after El-Moursy et al. 2001;
Abdel-Dayem 2004, 2012); Sabkhat al Shic (after Nussbaum 1987; Gebert 1991).

Genus Cicindela Linnaeus, 1758

Cicindela (s. str.) javeti azari Deuve, 2011

General distribution. Asia - Lebanon, Israel, Syria.

References. IsraEL — Valdenberg 1983: 42, 48 (as Cicindela campestris herbacea),
1985: 37 (as Cicindela campestris herbacea); Nussbaum 1987: 7-8 (as Cicindela herba-
cea); Wiesner 1992: 127 (as Cicindela herbacea); Puchkov and Matalin 2003: 105 (as
Cicindela herbacea); Rittner 2003 (as Lophyridia herbacea); Chikatunov et al. 2006:
293 (as Cicindela herbacea); Franzen 2007: 13 (as Cicindela herbacea); Ptashkovsky
2009: 8-9 (as Lophyra herbacea).

Distribution (Fig. 4). IsraEL, Upper Galilee: 1/z. Meron, 8.1V.1972, leg. D. Ger-
ling - 12 (TAU); Golan Heights: Mezudat Nimrod, 8.V.1983, leg. E. Shney-Dor
- 343 19 (TAU); Mt. Hermon: 1900 m, 22.IV.1973, leg. D. Furth - 14; 2000 m,
9.V1.1992, leg. A. Freidberg - 1¢ 19 (all TAU).

The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 127

LEBANON, Liban-Sud: Jezzin - 544 1199 (after Deuve 2011).
Syria, Dimashq: Bloudan (after Avgin and Wiesner 2009 - as Cicindela thughurica
Franzen, 2007).

(0) Cicindela (s. str.) herbacea herbacea Klug, 1832

General distribution. Asia - Lebanon.

References. LEBANON — Wiesner 1992: 127; Puchkov and Matalin 2003: 105;
Franzen 2007: 13; Deuve 2011: 129.

Distribution. LeBaNon, Liban-Nord: Bcharré, Les Cédres, VI. 1997 - 1419
(cIOv); Beharré - 19 (after Franzen 2007); Tannourine - 19 (after Deuve 2011).

Comments. Until recently both these species were recorded from Syria, Lebanon
and Israel by several authors as Cicindela herbacea Klug (Valdenberg 1983; Nuss-
baum 1987; Wiesner 1992; Puchkov and Matalin 2003; Chikatunov et al. 2006;
Franzen 2007; Ptashkovsky 2009). However, according to recent data C. herbacea
does not occur in Israel (Deuve 2011, 2012). The nominative subspecies occurs in
Lebanon and Syria; C. 4. aleppensis Deuve, 2012 is recorded from north-western
Syria, while C. 4. perreaui Deuve, 1987 and C. 4. colasi Deuve, 2011 are found in
Turkey - Tunceli and Adana Provinces, respectively. On the basis of the shape of
pronotum (Figs 38 vs 39), white elytral pattern (Figs 54 vs 55), size of aedeagus
and shape of it apex (Figs 93 vs 97), as well as shape of internal sack (Figs 94-96 us
98-100) we consider all studied specimens from Israel to be Cicindela javeti azari
Deuve, 2011 (type locality - Lebanon, Jezzine). It should be noted that the taxon-
omy of intraspecific forms within the ‘campestris-group is complex, and additional
studies are necessary.

Genus Cylindera Westwood, 1831

Cylindera (Eugrapha) contorta valdenbergi (Mandl, 1981)

General distribution. Asia — Israel, Egypt.

References. IsraEL — Bodenheimer 1937: 108 (as Cicindela); Mandl 1981: 169 (as
Cicindela); Naviaux 1983: 79; Valdenberg 1983: 43, 48 (as Cicindela), 1985: 29-30 (as
Cicindela); Nussbaum 1987: 7, 10 (as Cicindela); Werner 1992: 22, 48, 74; Wiesner
1992: 195 (as Cicindina); Puchkov and Matalin 2003: 110; Rittner 2003 (as Lophy-
ridia); Ptashkovsky 2009: 8-9 (as Lophyridia).

Distribution (Fig. 4). IsrazL, Northern Coastal Plain: ‘Akko, 7.VIII.1980,
leg. A. Valdenberg - 399; ‘Adlit, 5.VII.1942, B. Feldman - 14; Ma agan Mikha ‘el,
13.VII.1977, 9.1X.1978, 2.V.1979, 26.III.1980, VI.1980, leg. A. Valdenberg - 270.3
539 Q; 27.VI1.1979, leg. J. Kugler - 23°; 3.V1.1983, leg. E. Shney-Dor - 2¢5 792
(all TAU); VII.1987, leg. Y. Nussbaum — 19 (SIZ); 17.V.1980, leg. R. Naviaux - 19;

128 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

16.V.1986, leg. Y. Nussbaum - 53'¢ 699 (both cJW); Emeg Zevulun (after Nuss-
baum 1987). Central Coastal Plain: Bat Yam, 13.V11.1945, leg. H. Bytinski-Salz -
13 499 (TAU); Qesariya, Zerufa [Tsrufa] (both after Nussbaum 1987).

Genus Habrodera Motschulsky, 1862

(0) Habrodera nilotica nilotica (Dejean, 1825)

General distribution. Asia - Egypt (Sinai); Africa - Canary Islands (Grand Canary),
Senegal, Ghana, Mali, Guinea, Equatorial Guinea, Sierra Leone, Nigeria, Central Af-
rican Republic, Togo, Benin, Sudan, Egypt, Kenya, Congo, Zaire, Tanzania, Ethiopia,
Malawi, Mozambique, South Africa.

References. IsraEL — Chikatunov et al 2006: 293; Ecypr (Sina) - Alfieri 1976:
2 (as Cicindela); Wiesner 1992: 165); Werner 2000: 138; Abdel-Dayem et al. 2003:
202; Puchkov and Matalin 2003: 103; Abdel-Dayem 2004: 74.

Distribution (Fig. 5). Ecyrr (Sinat), Sinai Mountains: Wadi Isla (after Alfieri
1976; Abdel-Dayem et al. 2003; Abdel-Dayem 2004); St. Katherine (after Abdel-
Dayem 2004).

Comments. Previously Habrodera nilotica nilotica (Dejean, 1825) was mistakenly
referenced in the fauna of Israel (Chikatunov et al. 2006).

Genus Hypaetha LeCoute, 1857
(0) Hypaetha singularis (Chaudoir, 1876)

General distribution. Asia - Egypt (Sinai), Oman, Yemen; Africa - Egypt, Sudan,
Somalia, Eritrea, Djibouti.

References. Ecypt (Stnat) - Valdenberg 1983: 43, 45 (as Cicindela), 1985: 37 (as
Cicindela); Nussbaum 1987: 11, 13 (as Cicindela); Wiesner 1992: 219; Puchkov and
Matalin 2003: 112.

Distribution (Fig. 9). Ecyrr (Stnat), Southwestern Sinai: Nabeq, 8.V.1980, leg.
A. Valdenberg - 14; Ras Muhammad, 16.VII1.1978, leg. A. Valdenberg - 204 222
(all TAU).

Genus Lophyra Motschulsky, 1859
Lophyra (s. str.) flexuosa flexuosa (Fabricius, 1787)

General distribution. Europe - Portugal, Spain, Andorra, France, Italy, Switzerland;

Asia - Israel, Egypt (Sinai); Africa - Morocco, Tunisia, Algeria, Libya, Egypt.

The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 129

References. IsraEL — Bodenheimer 1937: 108 (as Cicindela), Valdenberg 1983:
42, 48 (as Cicindela flexosa), 1985: 33 (as Cicindela flexosa); Nussbaum 1987: 9, 15
(as Cicindela); Wiesner 1992: 160; Puchkov and Matalin 2003: 112; Chikatunov et
al. 2006: 293; Ptashkovsky 2009: 8-9; Ecypr (Stat) - Alfieri 1976: 1-2 (as Cicindela);
Nussbaum 1987: 9, 15 (as Cicindela); Wiesner 1992: 160; Abdel-Dayem et al. 2003:
203; Puchkov and Matalin 2003: 103; Abdel-Dayem 2004: 74.

Distribution (Figs 6, 9). IsRAEL (INCLUDING STATE OF PALESTINE), Northern
Coastal Plain: ‘Akko, 23.1V.1927, leg. O. Theodor - 19; Dor, 23.1V.1998, leg. A.
Traub - 299; Haifa, 18.V.1996, leg. Hauser - 19; Magan Mikhael, 16.V.1978,
24, X1.1978; 18:X11-1978, 12111979; 4:010.1979,, 10.10.1979 16.V1.1984, leg. A.
Valdenberg - 310.4 56929; 16.1V.1983, leg. E. Shney-Dor — 1¢ 399 (all TAU);
Central Coastal Plain: Bat Yam, 14.111.1940, 12.11.1941, leg. H. Bytinski-Salz -
2hS 19; Hofit, 21.1X.1994, leg. A. Freidberg - 14; Holon, 4.V.1978, leg. A. Frei-
dberg - 344 299; Nahal Alexander, 32°24'N, 34°52'E, 15.V.2005, leg. I. Zonstein
- 12; Rehovot, 18.11.1954, leg. J. Wahrman - 22° (all TAU); Southern Coastal
Plain: Ashdod, sands, 29.11.1984, leg. A. Freidberg - 14; Nir ‘Am, 21.1I1.1946, leg.
H. Bytinski-Salz - 2¢¢ 399; Nizzanim, 5.111.1996, leg. A. Freidberg - 19; Yavne,
17.IV.1974, leg. D. Furth - 243; Ziggim, 711.1996, leg. I. Yarom & A. Freidberg -
23¢ (all TAU); Dead Sea Area: Qumeran, 18.11.1997, leg. V. Chikatunov - 200;
Yeriho, 23.V11.2002, leg. V. Kravchenko & V. Chikatunov - 1¢ 299 (all TAU); Ara-
va Valley: ‘En ‘ddan, 18.1V.1999, leg. I. Yarom & V. Kravchenko - 2¢'¢ 19 (TAU);
Northern Negev: Beer Sheva, 1.1V.1946, leg. H. Bytinski-Salz - 2929; Bor Mashash,
18.IV.1995, leg. A. Freidberg - 14 19; Gevulot, 6.I1V.1985, leg. E. Shney-Dor - 3d
529; Revivim, Park Golda, 26.III.1999, leg. A. Freidberg - 1¢ 19 (all TAU); Cen-
tral Negev: Yeroham, 27.11.1966, leg. H. Bytinski-Salz - 4¢¢ 799; 25.1V.1973,
22.1V.1981, leg. J. Kugler - 1¢ 499, 19.11.1978, leg. M. Kaplan - 655 599;
19.III.1978, leg. A. Freidberg - 6334; Sede Boker, 8.V1.1987, leg. E. Shney-Dor — 13;
Kadesh Barnea, 11 1V.1974, leg. D. Furth - 344 29 9; 9.V.1979, leg. A. Valdenberg -
33¢ 629 (all TAU); Southern Negev: Elat, 14.V1.1981, leg. R. Keian - 19 (TAU).

Ecyrt (Srinat), Northern Sinai: Sabkhat al Shic, 8.V.1981, leg. A. Valdenberg
- 2464 299 (TAU); Lsmailia (after Alfieri 1976); Central Sinai Foothills: Wadi
Godirate |Gudeirat] (after Nussbaum 1987: Fig. 6; Abdel-Dayem 2004); Southwest-
ern Sinai: Nabeq (after Abdel-Dayem 2004), Wadi Gharandal (after Alfieri 1976;
Abdel-Dayem et al. 2003).

Genus Myriochila Motschulsky, 1858
Myriochila (s. str.) melancholica melancholica (Fabricius, 1798)
General distribution. Europe - Portugal, Spain, France, Italy, Malta, Albania, Greece,

Georgia, Armenia, Azerbaijan; Asia - Cyprus, Turkey, Lebanon, Israel, Egypt (Sinai),
Syria, Jordan, Saudi Arabia, Arab Emirates, Oman, Yemen, Bahrain, Kuwait, Iran,

130 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

Figure 6. Distribution of Lophyra flexuosa flexuosa in Israel and Palestine (open circles - records before
year 1949, half-solid circles - records between years 1950-1999, solid circles - records after year 2000; map
source - Eric Gaba Wikimedia Commons user: Sting and Wikimedia Commons user: NordNordWest,
URL - https://upload.wikimedia.org/wikipedia/commons/7/7c/Israel_relief_location_map.jpg).

The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 131

Iraq, Kazakhstan, Kyrgyzstan, Tadzhikistan, Uzbekistan, Turkmenistan, Afghanistan,
Pakistan (Punjab, Sind), India (Punjab, Haryana, Uttar Pradesh, Rajasthan, Madhya
Pradesh, Maharashtra, Bihar, West Bengal), China (Xinjiang); Africa - Cape Verde
Islands, Senegal, Morocco, Tunisia, Algeria, Libya, Egypt, Sierra Leone, Guinea, Guin-
ea Bissau, Chad, Ivory Coast, Togo, Ghana, Nigeria, Cameroun, Equatorial Guinea,
Central African Republic, Congo, Zaire, Kenya, Somalia, Ethiopia, Sudan, Tanzania,
Malawi, Mozambique, Angola, Namibia, South Africa, Madagascar, Seychelles.

References. IsraEL - Bodenheimer 1937: 108 (as Cicindela); Valdenberg 1983:
43, 46 (as Cicindela), 1985: 40 (as Cicindela); Nussbaum 1987: 9, 14 (as Cicin-
dela); Wiesner 1992: 211; Finkel et al. 2002: 28; Puchkov and Matalin 2003: 114;
Rittner 2003; Chikatunov et al. 2006: 293; Ptashkovsky 2009: 8-9; Ecypt (SINaI) -
Nussbaum 1987: 9, 14 (as Cicindela); Abdel-Dayem et al. 2003: 208; Abdel-Dayem
2004: 75.

Distribution (Figs 7, 9). ISRAEL (INCLUDING STATE OF PALESTINE), Upper Galilee:
Tel Dan, 25.V1I1.1958, leg. J. Wahrman - 14; 23.VIII.2002, leg. V. Kravchenko &
V. Chikatunov - 204; Nahal Keziv, 28.1X.1999, leg. M. Finkel - 14 19; Kefar Szold,
5.V.1998, leg. R. Ortal - 14; Hula, 1.V1.1968, leg. H. Bytinski-Salz - 655 328 all
TAU); Lower Galilee: Teverya, 3.V1.1961, leg. J. Wahrman - 443 39 9; 24.V.1981,
leg. A. Valdenberg - 34:5 19; Deganya, 15.1X.1951, J. Wahrman - 35d (all TAU);
Golan Heights: Hammat Gader, 23.VU.2002, 2.X.2002, leg. V. Kravchenko &
V. Chikatunov - 5¢¢ 499 (TAU); Northern Coastal Plain: Maagan Mikha ‘el,
17.V1.1973, leg. D. Furth - 5¢¢ 79.9; 20.1V.1986, leg. A. Freidberg - 20. ¢ 399 (all
TAU); Central Coastal Plain: Herzliyya, 20.V.2000, A. Freidberg - 299; Bet Dagan,
26.VIIL.1981, leg. Q. Argaman - 23°¢ 399; Ramat Gan, 3.V1.1985, leg. D. Gerling
-73¢5 529; Rosh Ha’Ayin, 15.X.1994, leg. V. Chikatunov - 3¢¢ 49.9; Tel Aviv,
2.1X.1974, leg. A. Freidberg & M. Kaplan - 5735'¢ 609 Q; 15.VII.2002, 12.IV.2003,
leg. V. Kravchenko & V. Chikatunov - 233 729; 24.VII.1948, H. Bytinski-Salz -
1Q (all TAU); Southern Coastal Plain: Nizzanim, 23.VUI.2002, 5.X.2002, leg. V.
Kravchenko & V. Chikatunov -4¢'4¢ 49 9 (TAU); Judean Desert: Nahal Perat (Wadi
Qelt), 23.V1I.2002, leg. V. Kravchenko & V. Chikatunov - 1d 299 (TAU); Jordan
Valley: Afigim, 26.V1I.1971, leg. M. Kaplan - 246; Maoz Hayyim, 21.V.1977, leg.
A. Valdenberg - 43° ¢ 29 @ (all TAU), from Dan to Ne‘ ot HaKikkar (after Nussbaum
1987); Dead Sea Area: Yeriho, 23.VU.2002, 5.X.2002, leg. V. Kravchenko & V. Chi-
katunov - 365 499; Qalya, 6.V.1980, leg. A. Valdenberg - 345 59 (all TAU);
Arava Valley: Gerofit, 2.VIII.2002 and 5.X.2002, leg. V. Kravchenko & V. Chikatu-
nov -23'¢ 499; Hazeva, 19.V11.1999, leg. I. Yarom & V. Kravchenko - 23: ¢ 32 9;
19.IX.1995, leg. A. Freidberg -19 19; En Tddan, 20.V1.1995, leg. A. Freidberg - 303
12; Yotvata, 24.VIII.1989, leg. A. Eitam - 10; Zugim, 22.V1.1999, leg. I. Yarom & V.
Kravchenko - 14 292.9; Samar, 29°50'N, 35°02'E, 26.IV.2007, leg. N. Ketner - 263
229 (all TAU); Northern Negev: Beer Sheva, 1.VIII.1945, leg. H, Bytinski-Salz -
244 299; Dimona, 18.VIII.1957, leg. J. Wahrman - 14 192; Gevulot, 18.V.1983,
6.V1.1984, 29. VIII.1987, leg. E. Shney-Dor - 11¢¢ 1299; Hazerim, 31.VIII.1951,
leg. J. Wahrman - 14 19; Retamim, 12.V1.2002, 5.VI1.2003, leg. V. Kravchenko &

A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

132

Figure 7. Distribution of Myriochila melancholica melancholica in Israel, Palestine and border areas of
Jordan (open circles - records before year 1949, half-solid circles - records between years 1950-1999, solid
circles - records after year 2000; map source - Eric Gaba Wikimedia Commons user: Sting and Wiki-
media Commons user: NordNordWest, URL - https://upload.wikimedia.org/wikipedia/commons/7/7c/

Israel_relief_location_map.jpg).

The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 133

V. Chikatunov - 204 599; Revivim, 1.1V.1942, leg. H, Bytinski-Salz - 245 299;
2.V1II.1958, leg. J. Kugler - 13 (all TAU); Ze'elim, 17.1X.1986, leg. Y. Nussbaum,
12 (cJW); Central Negev: Mash abbe Sade, 23.VUI1.1965. J. Wahrman - 330399;
27.VII1.1986, leg. A. Freidberg - 1¢ 19; Yeroham, 30.V.1957, leg. I. Yarkoni - 33
12; Tel Yeroham, 19.X1.1959, leg. L. Fishelsohn - 14; Ma’ agar Yeroham, 29.V11.2007,
leg. L. Friedman - 14; 30°59.37'N, 34°53.87'E, 22.V.2008, leg. L. Friedman - 263
22.9; Makhtesh Ramon, 9.VIII.1977, leg. A. Freidberg - 10; Mizpe Ramon, 4.VIII.1977,
leg. D. Simon - 14; Shivta, 23.V1.1978, leg. A. Freidberg - 535628 (all TAU); Qziot,
8.1X.1986, leg. Y. Nussbaum - 2¢'¢ (cJW); Ezuz (after Nussbaum 1987); Southern
Negev: Fiat, 6.[X.1974, leg. A. Freidberg - 243; Shizzafon, 12.V1.2002, 5.X.2001, leg.
V. Kravchenko & V. Chikatunov - 14 29 9 (all TAU).

Jordan, Al Balqa’: Al Maghtas, 23.V.1942, H. Bytinski-Salz - 19 (TAU).

Ecyrt (Sinat), Northern Sinai: £/ Arvish, 15.V1.1968, leg. J. Kugler - 1d 299
(TAU; including after Abdel-Dayem et al. 2003; Abdel-Dayem 2004); Southwestern
Sinai: Ofira, sewage, 2.V.1981, leg. A. Freidberg - 13 (TAU).

Tribe Megacephalini Laporte, 1834
Subtribe Megacephalina Laporte, 1834
Genus Grammognatha Motschulsky, 1850

Grammognatha euphratica euphratica Dejean in Latreille & Dejean, 1822

General distribution. Europe — Spain, Greece (Rhodes, Crete); Asia - Cyprus, Tur-
key, Lebanon, Israel, Jordan, Syria, Egypt (Sinai), Saudi Arabia, Arab Emirates, Ku-
wait, Oman, Yemen, Iran, Iraq, Pakistan; Africa - Morocco, Tunisia, Algeria, Libya,
Egypt, Djibouti.

References. IsraEL — Bodenheimer 1937: 108 (as Megacephala); Naviaux 1983:
75 (as Megacephala), Valdenberg 1983: 42, 47, 1985: 40 (as Megacephala); Nussbaum
1987: 8, 11 (as Megacephala); Wiesner 1992: 44 (as M. euphratica nigra); Franzen
2001: 89 (as Megacephala); Puchkov and Matalin 2003: 118 (as Megacephala); Rittner
2003 (as Megacephala); Chikatunov et al. 2006: 293 (as Megacephala); Ptashkovsky
2009: 8-9 (as Megacephala); Ecypr (Stal) - Schatzmayr 1936: 6 (as Megacephala);
Alfieri 1976: 1 (as Megacephala); Nussbaum 1987: 8, 11 (as Megacephala); Wiesner
1992: 44 (as Megacephala euphratica nigra); Werner 1999: 68 (as Megacephala); El-
Moursy et al. 2001: 66 (as Megacephala); Franzen 2001: 88 (as Megacephala); Abdel-
Dayem et al. 2003: 196; Puchkov and Matalin 2003: 118 (as Megacephala); Abdel-
Dayem 2004: 73.

Distribution (Figs 8, 9). IsRAEL (INCLUDING STATE OF PaLEsTINE), Northern
Coastal Plain: Haifa (after Franzen 2001); ‘Adit, 4.V1.1979, 1.V.1979, leg. A. Val-
denberg - 29 9; 4.V1.1983, leg. E. Shney-Dor - 14; 32°42'N, 34°56'E, 17.V.1997,
leg. E. Orbach - 1¢ 19 (all TAU), VI.1989, leg. E. Orbach - 1¢ (cJW); V.1989,
not far from the coastal line, running to light, leg. E. Orbach - 234 (after Werner

134 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

~ ‘asin

ede Lt abl

mye

Figure 8. Distribution of Grammognatha euphratica euphratica in Israel and Palestine (open circles -
records before year 1949, half-solid circles - records between years 1950-1999, solid circles - records
after year 2000; map source - Eric Gaba Wikimedia Commons user: Sting and Wikimedia Commons
user: NordNordWest, URL - https://upload.wikimedia.org/wikipedia/commons/7/7c/Israel_relief_loca-

tion_map.jpg).

the tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 135

Figure 9. Distribution of Hypaetha singularis (red circles), Myriochila melancholica melancholica (blue
thombs), Lophyra flexuosa flexuosa (orange squares) and Grammognatha euphratica euphratica (lilac ti-
angles) in Sinai Peninsula, Egypt (open symbols - records before year 1949, half-solid symbols — records
between years 1950-1999; URL map source - https://upload.wikimedia.org/wikipedia/commons/5/59/

Sinai_relief_location_map.svg).

136 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

1999); Dead Sea Area: Bet Ha’Arava, 5.1V.1941, leg. O. Theodor - 25'¢ 292 9; Jordan
River, near Dead Sea, 5.IV.1941, leg. O. Teodor - 12 (TAU); ‘En Gedi, 24.III.1958,
leg. J. Kugler - 244; 15.I01.65, leg. K. Yefenof - 12; Ne‘ot Hakikkar, 15.11.1999,
19.IV.1999, leg. I. Yarom & V. Kravchenko - 34.5 19; Qalya, 11.1V.1958, leg. M.
Pener - 2.44 19; Sedom, 6.V.1961, at night, leg. J. Wahrman - 13; Shefekh Zohar,
16.IV.1980, leg. J. Kugler - 19; Zomet Zohar, 17.1V.1997, leg. L. Friedman - 19
(all TAU); Nawit Pools (after Nussbaum 1987); Arava Valley: Gerofit, 23.1V.2003,
12.V.2003, 6.VI.2003, leg. D. Utshitel & V. Chikatunov - 34.4 29 2 (TAU); South-
ern Negev: Eat, 30.VIII.1959, leg. L. Fishelsohm — 1 3 (TAU).

Ecypt (Srnat), Northern Sinai: Sabkhat al Bardawil, 23.11.1969, leg. A. Nitzan
- 144 222 (TAU); Arish (after Abdel-Dayem 2004); Zaranik Protectorate (after
El-Moursy et al. 2001; Abdel-Dayem 2004); Sinai Mountains: Da/hab (after Nuss-
baum 1987); Southwestern Sinai: F/ Tor (after Schatzmayr 1936; Alfieri 1976;
Nussbaum 1987; Abdel-Dayem et al. 2003; Abdel-Dayem 2004); Nabeg (after
Nussbaum 1987).

An identification key to the tiger beetles of Israel and adjacent lands

1(2) Anterior angles of pronotum projected towards the margin of prothorax
(Figs 10, 45); the fourth joint of maxillary palpus shorter than the third
one...... Megacephalini (Grammognatha euphratica euphratica Dejean, 1822)

2A) Anterior angles of pronotum not projected towards the margin of prothorax
(Figs 12-14); the fourth joint of maxillary palpus longer than the third one...
Bolen ete eee ee XA 2 Ree) AEE eR Cicindelini 3
3(4) Proepisterna prominent on pronotum so pronotopleural suture clearly visible

dorsally (Fig. 42); anterior margin of pronotum with row of flat white setae ....

sei dette eect ttt Hypaetha ((°) Hypaetha singularis (Chaudoir, 1876))

4(3) Proepisterna not prominent on pronotum so pronotopleural suture not visible
dorsally (Figs 30-41, 43-44); anterior margin of pronotum glabrous........... 5

5(6) Labrum with four submarginal setae (Fig. 29); middle and hind femora

with numerous hooked setae along posterior margin, hind femora with

sparse-hooked Sete tic. G4) Foi, t-te cusses siesta tasatnys duct tbaashcnsae Myriochila
(s. str.) (Myriochila (s. str.) melancholica melancholica (Fabricius, 1798))
6(5) Labrum with at least six submarginal setae, except aberrant specimens with
3—5 setae (Figs 15-26, 28); femora without hooked setae along posterior
IVIATPAN vartacauw gedauekctanaeedidaakonsdaccewt tabh cus eauees gaat kasashertuaseng peace gh eal ela Th
7(8) Genes pilose (bigs) UG, gi not wees creed See toe ee, Se Re 9
8(7) OPO aE TeeSdbtel cosine esa bese [io ) EAR ORR A an GI aa ee OI yl a CN 13
9(10) Clypeus glabrous, anterior and posterior margins of each eye with group of

white decumbent setae; labrum with 10 submarginal setae in a single row
(Fig. 25); fourth antennomere of males with penicillus (Fig. 14); white
elytral pattern with complete humeral lunule, long sinuate middle band

10(9)

11(12)

12(11)

12a(12b)

12b(12a)

12c(12d)

12d(12c)

13(14)

14(13)

The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 137

and apical lunule coupling together via marginal and sutural bands (Fig.
SG) RG outirritae te Habrodera ((°) Habrodera nilotica nilotica (Dejean, 1825))
Clypeus pilose, anterior and posterior margins of each eye glabrous; labrum
with several rows of numerous submarginal setae (Figs 15-18); fourth an-
tennomere of males glabrous (Fig. 11); white elytral pattern without mar-
ginal and sutural bands (Figs 46-49) ooo. eeeeseeseeereereeeees Calomera 11
Elytra dark brown with purple-bronze or green reflection (Fig. 46); pronotum
1.05—1.15 times as wide as long with straight parallel or slightly convergent
lateral sides (Fig. 30); aedeagus straight, with long thin basal portion, apical
lobe with distinct lateral flanges and small hook, without central groove (Figs
73, 77), ventro-apical bladder of internal sac short, right and left basi-lateral
bladders very large (Figs 77, 81)... Calomera aulica aulica (Dejean, 1831)
Elytra green sometimes with bronze or blue reflection (Figs 47-49); pro-
notum 1.15—1.35 times as wide as long with rounded distinctly conver-
gent lateral sides (Figs 31-33); aedeagus curved, with short basal por-
tion, apical lobe without lateral flanges and hook, but with clear central
groove (Figs 74-76, 82-84), ventro-apical bladder of internal long, right
and left basi-lateral bladders as small acicular areas (Figs 78-80, 82-
ZB ees co ESE Lk UNC Calomera littoralis (Fabricius, 1787) 12a
Left mandible with four teeth distal to apical molar (Fig. 16); pronotum
narrow, 1.15—1.2 times wider than long (Fig. 31); aedeagus with small dis-
tinct bulge on the dorsal surface (Fig. 74); ventro-apical bladder of internal
sac long and curved towards and on the left, apex of medial tooth blunt
(Figs 74, 78, 82)... Calomera littoralis aulicoides (J.R. Sahlberg, 1913)
Left mandible with three teeth distal to apical molar (Fig. 17-18); prono-
tum wide, 1.2—1.35 times wider than long (Figs 32-33); aedeagus without
bulge on the dorsal surface (Figs 75—76); ventro-apical bladder of internal
sac straight and not curved, apex of medial tooth sharp (Figs 75-76, 79-80,
BOS OA un Ael eas: air S00 or sya nina Aer maui setae 0. East, tanita td 2 etter tn ene oe 12c
Labrum wider, 2.6—2.65 times as wide as long (Fig. 32), base of medial
tooth of internal sac with one rarely two small additional spikes (Figs 75, 79,
BGI 2 wits Crees eee ee. oF Rava Calomera littoralis winkleri (Mandl, 1934)
Labrum narrower, 2.35—2.45 times as wide as long (Fig. 33), base of me-
dial tooth of internal sac smooth, without additional spikes (Figs 76, 80,
Arsene Mati! ae vac os (0) Calomera littoralis nemoralis (Olivier, 1790)
Labrum tridentate with distinctly prominent apical teeth; mandibles with
two teeth distal to apical molar (Fig. 26); scapus covered by numerous white
decumbent setae (Figs 13, 26), fourth antennomere of males with penicil-
lus (Fig. 13); posterior margin of each eye with group of white decumbent
setae; white elytral pattern with basal dot and incomplete sutural band (Fig.
57)....Lophyra (s. str.) (Lophyra (s. str.) flexuosa flexuosa (Fabricius, 1787))
Labrum unidentate (Fig. 20-24, 28), in some species tridentate but with not
or slightly prominent apical teeth only (Fig. 19); mandibles with three teeth

138

15(16)

16(15)

17(18)

18(17)

19(20)

20(19)

21(22)

22(21)

23(24)

A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

distal to apical molar (Fig. 19-24); scapus glabrous (Figs 19-22, 28) or only
with several sparse setae except apical ones (Figs 23-24), fourth antennomere
of males glabrous (Fig. 11); posterior margin of each eye glabrous; white ely-
tral pattern without basal dots and sutural band (Figs 50-55, 59) ......... 15
Head glabrous; scapus with apical setae only (Fig. 19-22, 28); lateral side of
pronotum pilose (Figs 34-37, 43); white elytral pattern with long marginal
band and long sinuate middle band (Figs 50-53, 59) ....sceceseseeseeseereees 17
Frons and vertex with long soft hairs, scapus with several setae except api-
cal ones (Figs 23-24); lateral side of pronotum with soft sparse setae (Figs
38-39); white elytral pattern without marginal band and only with short

slightly curved middle band (Figs 54-55) oo... eee Cicindela (s. str.) 25
Anterior portion of apical lunule long, extending basal transverse portion of
middle baid"(Big< 59 )eeater eter ests rotte cates caetttae foe ede arose Cylindera

(Eugrahpa) (Cylindera (Eugrapha) contorta valdenbergi (Mandl, 1981))
Anterior portion of apical lunule short, extending only apical portion of
iniddle band (Pigs 50-53) e.cesescsecccrsuccnesdenssecesenre Cephalota (Taenidia) 19
Labrum tridentate, relatively short, no less than 2.3 times as wide as long (Fig.
19); pronotum 1.2—1.4 times wider than long (Fig. 34); mesepisternum en-
tirely covered by white setae, densely in males and sparsely in females; white
elytral pattern with relatively broad marginal band coupling with humeral and
apical lunule as well as with middle band (Fig. 50), apical margin of elytra
in sexes wide rounded, subtend practically right angle with sutural tooth
(Figs 65-66); aedeagus with long thin gradually curved basal portion (Fig.
3) ee OP (0) Cephalota (Taenidia) tibialis tibialis (Dejean, 1822)
Labrum unidentate, relatively long, no more than 2.3 times as wide as long
(Figs 20-22); pronotum 1.1—1.25 times wider than long (Figs 35-37);
mesepisternum covered by white setae only along posterior margin and
on the base; white elytral pattern usually with narrow marginal band or
without it so in some specimens humeral lunule distinctly separated (Figs
51-53), apical margin of elytra at least in males subtend acute angle with
sutural tooth (Figs 67-72); aedeagus with short thin basal portion (Figs 87,

4-11" antennomeres dark brown; elytra bright purple, 1.5—1.6 times as long as
wide (Fig. 51), apical elytral margin in females narrowly rounded and subtend
small right angle with sutural tooth (Fig. 67-68); aedeagus with broad blunt apex
(Figs 87-88) ......... (0) Cephalota (Taenidia) littorea littorea (Forskal, 1775)
4-11" antennomeres light brown or yellowish; elytra greenish or greenish-
blue sometimes with distinct golden-purple reflection, no less than 1.65
times as long as wide (Figs 52-53), apical elytral margin in both sexes sub-
tend acute angle with sutural tooth (Figs 69-72); aedeagus with arrow-
SHAD ECa eK VUE1G6 B29 Des Ft ecanwrde aps ocesenc a cose vectes sev cs save pant gue tonclcee oai'ent 23
Labrum shorter, 2.0—2.2 times as wide as long (Fig. 21); lateral side of prono-
tum straight, slightly convergent to large posterior angles (Fig. 36); humeral

The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 139

lunule separated or narrowly coupled with marginal band (Fig. 52); aedeagus
larger, with relatively long thin basal portion (Fig. 89) and short tapered apex
(Fig. 90) ... Cephalota (Taenidia) zarudniana vartianorum (Mandl, 1967)
24(23) Labrum longer, 1.6—1.7 times as wide as long (Fig. 22); lateral side of prono-
tum slightly rounded, distinctly convergent to small posterior angles (Fig. 37);
humeral lunule coupled with middle band via marginal band (Fig. 52); aedea-
gus smaller, with short thin basal portion (Fig. 91) and long tapered apex (Fig.
92).....(0) Cephalota (Taenidia) deserticola deserticola (Faldermann, 1836)
25(26) Pronotum with convex lateral sides gradually convergent to posterior angles,
anterior margin same length or slightly longer than posterior one, notopleural
suture looks like smooth border (Fig. 39); mesepisternum of female with small
shallow pit and deep all along coupling sulcus, mesepimeron with groove along
anterior margin (Fig. 63); middle band of white elytral pattern without oblique
strip between transverse basal and circled apical portions, basal portion of api-
cal lunule small (Fig. 55); aedeagus shorter, no more than 0.55 times as long
as elytra (Fig. 97); basal and right ventro-lateral bladders of internal sac short
GEER SSO) et catenentsenanamedtess. Cicindela (s. str.) javeti azari Deuve, 2011
26(25) Pronotum with straight lateral sides sharply convergent to posterior an-
gles, anterior margin clearly longer than posterior one, notopleural su-
ture looks like cut border (Fig. 38); mesepisternum of female with deep
apically but shallow and indistinct basally coupling sulcus only, mese-
pimeron without groove along anterior margin (Fig. 62); middle band
of white elytral pattern with distinct oblique strip between transverse
basal and circled apical portions, basal portion of apical lunule large
(Fig. 54); aedeagus longer, no less than 0.6 times as long as elytra (Fig.
93); basal and right ventro-lateral bladders of internal sac long (Figs 94—
BTC s bead, RUAN ER ee (0) Cicindela (s. str.) herbacea herbacea Klug, 1832

Distribution

With these current records, eight species of tiger beetles, one of them with two sub-
species, belonging to seven genera of two tribes are known from Israel (Table 1). The
Rift Valley, including Jordan Valley, Dead Sea area and Arava Valley, with six cicinde-
lids species is the most speciose region. The Coastal Plain is the second richest region
with five species. The species richness gradually decreases from Northern (fife species)
through Central (four species) to Southern (three species) Coastal Plain. In the central
densely populated areas of Israel, such as Samaria and Judea, the least number of tiger
beetles species are recorded. Among all MZ. melancholica melancholica is the most com-
mon species observed in all regions of the country (Table 1, Fig. 7), while L. flexuosa
flexuosa is the second most widespread species of tiger beetles absent only from north-
ern (Galilee, Golan Heights) and central (Samaria, Judea) regions (Table 1, Fig. 6).
Lophyra flexuosa (Fabricius, 1787) reaches the eastern limit of its distribution in Israel.

A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

140

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The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 141

i i | NN a
Ge =

Figures 15-22. Labrum and mandibles of males, dorsal view: 15 Calomera aulica aulica \6 Calomera
littoralis aulicoides \1 Calomera littoralis winkleri \8 Calomera littoralis nemoralis \9 Cephalota tibialis
tibialis 20 Cephalota littorea littorea 2\ Cephalota zarudniana vartianorum 22 Cephalota deserticola deser-

ticola. Scale bars: 1 mm.

142 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

Figures 23-29. Labrum and mandibles of males, dorsal view: 23 Cicindela herbacea herbacea 24 Cicin-
dela javeti azari 25 Habrodera nilotica nilotica 26 Lophyra flexuosa flexuosa 27 Hypaetha singularis 28 Cyl-
indera contorta valdenbergi 29 Myriochila melancholica melancholica. Scale bars: 1 mm.

143

the tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands

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144

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the tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 145

71 12

Figures 62-72. Details of Cicindelinae: 62-63 mesoepisternal coupling sulcus 64 hind femora 65-72 api-
cal part of left elytron 62 Cicindela herbacea herbacea 63 Cicindela javeti azari 64 Myriochila melancholica
melancholica 65-66 Cephalota tibialis tibialis 61-68 Cephalota littorea littorea 69-10 Cephalota zarudniana
vartianorum T1=12. Cephalota deserticola deserticola 64-65, 67, 69, 71 males 62-63, 66, 68, 70, 72 fe-
males. Scale bars: 1 mm (62-63: A; 64: B; 65-66: C; 67-68 D; 69-70: E; 71-72: F).

A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

146

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the tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 147

Figures 77-84. Internal sack of Calomera spp.: 17, 81 C. aulica aulica 18, 82 C. littoralis aulicoides
79, 83 C. littoralis winkleri 80, 84 C. littoralis nemoralis 77-80 right lateral view 81-84 dorsal view;
BLR - basi-lateral right bladder; BLL - basi-lateral left bladder; VA — ventro-apical bladder; mt — median

tooth. Scale bar: 1 mm.

A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

148

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the tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 149

Cc

Figures 93-100. Aecdeagus and internal sack of Cicindela ssp.: 93-96 C. herbacea herbacea 91-100 C.
javeti azari 93, 97 aedeagus 94-96, 98-100 internal sac 93, 96-97, 100 left lateral view 95, 99 dorsal
view 94, 98 right lateral view 98-100 partly inflanted); B - basal bladder; VZR — ventro-lateral right
bladder. Scale bars: 1 mm (93, 97: A; 94—96: B; 98-100: C).

Three subspecies, C. contorta valdenbergi, C. javeti azari and C. zarudniana var-
tianorum, are characterized by a restricted distribution in Israel (Fig. 4). The first two
first subspecies should be considered as regional endemics.

The nominative subspecies of Cylindera contorta (F.-W., 1828) is widely distrib-
uted in Central Asia, some regions of Cis- and Transcaucasia as well as in the northern
and western sides of the Black Sea from southern Russia to Romania (Wiesner 1992;
Cassola 1999; Puchkov and Matalin 2003), however it is not known from Anatolia
(Corel 1988; Cassola 1999; Puchkov and Matalin 2003; Avgin and Ozdikmen 2007),
Syria (Wiesner 1992; Puchkov and Matalin 2003; Avgin and Wiesner 2009; Jaskula
and Rewicz 2014), Jordan (Wiesner 1992; Puchkov and Matalin 2003), Iraq (Ali 1978;
Wiesner 1992; Puchkov and Matalin 2003) and Saudi Arabia (Wiesner 1992; Cassola
and Schneider 1997; Puchkov and Matalin 2003; Al Ahmadi and Salem 1999). The
populations of C. contorta valdenbergi inhabit the Mediterranean coast from ‘Akko
(Northern Coastal Plain) to Bat Yam (Central Coastal Plain) in Israel (Nussbaum
1987; our data) as well as between Ras El Bar and Abu Qir in north-eastern Egypt
(Alfieri 1976; Abdel-Dayem et al. 2003) are distinctly scattered and bound the south-
western limit of the distributional area of C. contorta as a whole.

A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

150

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The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 151

Cicindela javeti azari has a restricted distributional area and now is known only
from southern Lebanon (Deuve 2011), southwestern Syria (Avgin and Wiesner 2009)
as well as northern regions of Israel: Upper Galilee and Golan Heights (Nussbaum
1987; our data). Among three known subspecies (Deuve 2011) C. javeti azari inhabits
the southern part of the species range area.

Cephalota zarudniana vartianorum lives from south-eastern Iran across Iraq and
Syria to Jordan and Israel (Wiesner 1992; Puchkov and Matalin 2003). The Dead Sea
Area is the western border of the distributional area both for this subspecies as well as
for the species as a whole.

It should be noted that the three mentioned above subspecies were recorded in
Israel only during XX century (Fig. 4), and the latest records are dated from the late
80’s to the early 90’s.

The Sinai Peninsula is the most diversity of tiger beetles region from all neighbou-
ring territories by Israel because nine species live here, and C. tibialis tibialis, C. littorea
littorea, H. singularis and H. nilotica nilotica are never really observe in Israel (vs Chikatu-
nov et al. 2006). Among them C. tibialis tibialis is an endemic of Egypt and occurs along
Mediterranean Sea coast in the Governorates Matrouh, Alexandria, Kafr el-Sheikh,
Damietta, Port Said and North Sinai (Gebert 1991; Abdel-Dayem et al. 2003; Abdel-
Dayem 2012). Moreover, C. /ittorea littorea is an regional near-endemic living along Red
Sea coast in Egypt and Saudi Arabia (Gebert 1991; Cassola and Schneider 1997; Abdel-
Dayem et al. 2003). Arabian-African Hypaetha singularis lives along Red Sea coast in
Egypt, Sudan, Eritrea and Yemen, and on the shore of Gulf of Aden in Djibouti, Somalia
and Yemen (Wranik et al. 1991; Werner 2000; Wiesner 2002, 2005) as well as on the lit-
toral of Arabian Sea in Oman (Cassola and Rihane 1996). The Sinai localities are limited
the northern border of the distribution area of this species. African Habrodera nilotica
nilotica is widely distributed in Afrotropical Region (Wiesner 1992; Werner 2000). Two
known localities from Sinai Mountains (Alfieri 1976; Abdel-Dayem et al. 2003; Abdel-
Dayem 2004) are limited the distribution range of this species to the east.

According to the analysis of the similarity between faunas of tiger beetle of natural
regions of Israel and the Sinai Peninsula two large clusters are recognized (Fig. 101).
First of them includes the faunas associated with southern part of the Great Rift Valley
(Arava valley and Dead Sea area) and most part of the Sinai Peninsula, while the second
combine most Israeli regions as well as Central Sinai Foothills. The last cluster diverges
on the four groups. The fist combines assemblages of tiger beetles of the Mediterranean
coastal habitats within the Northern and Central Coastal Plains. The communities
typical for the arid habitats of the Negev Desert and the Central Sinai Foothills as
well as for coastal habitats of the Southern Coastal Plain form the second group. The
third group includes assemblages of the northern not seashore habitats of the Jordan
Valley, Lower Galilee and Golan Heights. The last group is artificial, because the fauna
of tiger beetles of Judea should be most similar to the fauna of the Dead Sea Area or
the Northern Negev, while the fauna of tiger beetles of the Galilee, Jordan Valley and
Golan Heights should be the most similar to each other. First of all, this discrepancy is
due to a lack of data about tiger beetles of the central regions of Israel.

152 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

Uper Galilee

Judean Desert

Judean Hills

Lower Galilee

Golan Heights

Jordan Valley
Southern Coastal Plain — =

Northern Negev

Central Negev

Southern Negev

Central Sinai Foothills

Northern Coastal Plain

Central Coastal Plain

Dead Sea Area

Arava Valley

Northen Sinai

Southwestern Sinai

Sinai Mountains

0 10 20 30 40 50 60 70 80 90 100
Similarity (%)
Figure 101. Similarities between tiger beetle faunas of different regions of Israel and the Sinai Peninsula

(Complete linkage procedure, squared Euclidean distances).

Phenology

According to the literature data (Alfieri 1975; Nussbaum 1987; Abdel-Dayem et al.
2003) and the results of our own study some aspects of the phenology of tiger beetles
both in Israel and on the Sinai Peninsula are discussed. The period of activity of the
beetles but not the breeding period was analysed first of all. As a result, five groups of
the tiger beetles were obtained (Table 2). Three species with the longer period of activ-
ity from January to November or from February to December belong to the all-year
group. Five species, including two subspecies of Calomera littoralis (F., 1787), charac-
terized by the prolonged period of activity from February to October-November, from
March-April to November or from March to December and form the richest spring-
fall group. Two species recorded only on the Sinai Peninsula with the period of activity
from May to August-September are composed the summer group. At last, both the
spring group (activity from February to May) and the spring-summer group (activity
from February to August) contain a single species each.

It should be noted that the period of activity of some studied species does not
correspond with the data of previous studies in Israel (Nussbaum 1987) and on the
Sinai Peninsula (Alfieri 1975; Abdel-Dayem et al. 2003), as well as in the other parts
of the distribution area (Jaskuta and Rewicz 2015; Jaskuta et al. 2015). For example,
the activity of C. aulica aulica, C. littoralis aulicoides, C. zarudniana vartianorum, C.
contorta valdenbergi and G. euphratica euphratica start one-two months earlier, while
the activity of C. aulica aulica, C. littoralis aulicoides, M. melancholica melancholica and
G. euphratica euphratica finish one-three, and in the case with L. flexuosa flexuosa even

The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 153

Table 2. The phenology of tiger beetles in Israel (grey — our data; pink — after Nussbaum 1987) and in
the Egypt (green — after Alfieri 1975; blue — after Abdel-Dayem et al. 2003).

Cephalota zarudniana &
vartianorum &
(Si) id
Grammognatha euphratica NES
ieee nS
iphratica (Si) AS
Si Si
Hypaetha singularis
Oye Law S
Si Sa | 357 S
7s imo eae le zeu lanyae lense a)
Cephalota littorea littorea (Si) (Si)
(Si)
Sian Stile
Cephalota tibialis tibialis al ey ae dee
S27 ESE Ses le Sea gee ESE. se" 572 peor
Calomera littoralis
aulicoides fs |
Calomera littoralis winkleri —
a
=
Cylindera ee
contorta valdenbergi
Cicindela javeti azari
Myriochila melancholica
melancholica
ttc tae Si
Habrodera nilotica nilotica
(Si) | (Si) (Si) | (Si)
; A
Calomera aulica aulica Si (Sil Si S
(Si) | (Si) | 0S2) | (S02) (Sd) | (Sd) | 652) | (S22) =
(Si)
Lophyra flexuosa flexuosa (Si) | (SD
I TT | el | BV) Mo OM eV eT}, EX |) |) XT | OXTT

Notes. Si — records only on the Sinai Peninsula, (S7) — records including the Sinai Peninsula. The density

of the grey color corresponds with the frequency of the records of species (subspecies):

1-3 4-6 7-9 10-12

154 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

six months later comparing with the data of Nussbaum (1987). On the other hand,
Nussbaum (1987) indicated longer period of activity of C. /ittoralis winkleri and C.
tibialis tibialis as well as the later finish of the activity of C. contorta valdenbergi and C.
javeti azari (Table 2).

Similarly, the periods of activity of C. aulica aulica, L. flexuosa flexuosa and M. mel-
ancholica melancholica in the central and southern Levant as well as on the Sinai Penin-
sula are appreciably longer than in the Maghreb region. So, in Tunisia C. aulica aulica
records only in June and July (Jaskuta and Rewicz 2015), while in Israel it active from
March to December and on the Sinai Peninsula from February to October (Table 2).
Both in Tunisia and Morocco the period of activity of L. flexuosa flexuosa lasts from
March-April to July (Jaskula and Rewicz 2015; Jaskuta et al. 2015) but in Israel it
continues from February to December (Table 2).

On the contrary, in Tunisia the activity of G. euphratica euphratica begins in March
and ends in July (Jaskuta and Rewicz 2015) that is similar with the period of activity
in Israel and on the Sinai Peninsula (Table 2), while in Morocco it takes only three
months - from June to August (Jaskula et al. 2015). The same situation is observed
for different subspecies of Cephalota littorea (Forskal, 1775) as well as C. Littoralis. In
Tunisia C. littorea gouditii (Dejean, 1829) is active from May to October (Jaskula and
Rewicz 2015) while the period of activity of C. /ittorea littorea on the Sinai Peninsula
lasts from May to September (Table 2). The activity of C. Littoralis littoralis in Morocco
is observed from April to October (Jaskuta et al. 2015) and in Tunisia from March to
August (Jaskula and Rewicz 2015), while the activity of C. littoralis aulicoides in \srael
and on the Sinai Peninsula as well as C. Littoralis winkleri in Israel occurs from Febru-
ary to October and from February to November, respectively (Table 2).

However, we must remember that the obtained data are compilative. The differ-
ences in the time and the density of sampling, the collection technics as well as the
frequency of visit of the particular localities and habitats could really distort the real
pattern.

Faunogenesis

The tiger beetle fauna of Israel as well as the Levant as a whole is complex. In geologi-
cal time these areas were settled by species from different Mediterranean, African and
Asiatic regions.

Unfortunately, the information about fossil Cicindelinae is extremely scant (Na-
gano et al. 1982). At present time South American Oxycheilopsis cretacicus Cassola &
Werner, 2004 (Lower Cretaceous ca. 125 Ma) is the oldest known fossil tiger beetle
(Cassola and Werner 2004). Three samples of fossil cicindelids are known from the
northern Europe Baltic Amber (Oligocene ca. 23-34 Ma). Despite the identification
ambiguity of the species, the genera were interpreted as the recent ones (Nagano et al.
1982; Réschmann 1999) as most known fossil Carabidae and other Coleoptera (Alek-
seev 2013). All other fossil records of the tiger beetles from the Europe and northern

The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands 155

America (USA and Canada) are dated from the Quaternary period from Pleistocene to
Holocene, and all other species are interpreted as recent (Nagano et al. 1982).

By analogy with other groups of carabid beetles (Kataev 1984, 2011; Casale and
Vigna Taglianti 1999; Ruiz et al. 2012), we can assume that the genesis of the ancestral
taxa of most recent cicindelids in the Mediterranean region began in late Paleogene —
early Neogene (on the border of Oligocene — Miocene). According to data of DNA
analysis the divergence processes of taxa of subtribe Cicindelina began ca. 15-25 Ma
with most intensity between 2-10 Ma (Barraclough and Vogler 2002; Pons et al.
2004; Tsuji et al. 2015). For example, the diversification of the species within Cicin-
dela hybrida group started ca. 2 Ma (Cardoso and Vogler 2005), while the separation
of the genus Cosmodela Rivalier, 1961 from other Cicindelinae took place ca. 2.2—5
Ma (Lopez-Lopez et al. 2015; Tsuji et al. 2015). Based on the fossil material we could
be argued that at least 60,000-—70,000 yrs. BP the recent species of tiger beetles were
already presented both in the North America and in the Eurasia (Nagano et al. 1982).

The continental drift of the Arabian and Anatolian Plates, their collision and, as the
result, closing the Neotethys Ocean during Oligocene-Miocene were the most important
processes forming the Mediterranean Sea and the genesis of the terrestrial Mediterranean
fauna. The Eurasian-African land-bridge formed during late Burdigalian — middle Ser-
ravallian ca. 12.5-18 Ma (Régl 1998) initiated the species change/exchange between the
Europe, Asia and Africa (Koufos et al. 2005). The territory of the Sinai Peninsula and the
Levant free from the sea formed the first transit corridor. However, it was interrupted
at least twice in Langhian (ca. 16—16.4 Ma) and in early Serravallian (ca. 13-13.3 Ma),
while in Tortonian (ca. 11.6 Ma) the final connection of Arabian and Anatolian plates
and isolation of the Mediterranean Sea took place (Régl 1998, 1999). Because the Cen-
tral and Southern Levant as well as the Sinai Peninsula were the part of the Arabian plate
connected with the African continent (Régl 1998; Popov et al. 2004; Robertson et al.
2012; Berra and Angiolini 2014) the African species G. euphratica, H. nilotica, M. mel-
ancholica and L. flexuosa could have colonized these territories before the other species.

The sharp decrease of the level of the Mediterranean Sea in Messinian (ca. 5.5—6
Ma) caused the formation of both numerous shallow enclosed saline basins and the
land-bridges between Southern Europe and Northern Africa (Rogl and Steininger
1983). In our opinion during this time the active divergence and dispersion of such
halophilic genera as Cephalota, Calomera and Hypaetha as well as the species of the
subgenus Eugrapha occurred. All of them are arisen in the saline landscapes along
the seashores of Para- and Neotethys in the Southern Russland as well as Central and
Western Asia. From these regions the ancestors of the recent taxa probably dispersed
through the Middle East, Arabian Peninsula and Anatolia to the Levant and the Sinai
Peninsula, and some of them to Northern Africa. The second stream of the migration
was possible along the Mediterranean coast of Southern Europe. Following this some
species reached the Iberian Peninsula, and then the western regions of Northern Af-
rica. In contrast G. euphratica, M. melancholica, L. flexuosa could be populated South-
ern Europe (Garcia-Reina et al. 2014), Western and Central Asia as well as Sind and
some regions of South-Eastern Asia. Finally, possible during the last Glacial Period,

156 A.V. Matalin & VI. Chikatunov / ZooKeys 578: 115-160 (2016)

the ancestors of C. javeti and C. herbacea dispersed into the Levant from the Anatolia,
a region characterized by a higher level of diversity of the species of the Cicindela camp-
estris group (Cassola 1999; Franzen 2007; Deuve 2011, 2012; our unpublished data).
This proposed version of the biogeographical genesis of the fauna of tiger beetles
of the Levant should be considered an initial hypothesis. Molecular analysis and more
detailed paleontologic information are necessary to robustly reject or validate it.

Acknowledgements

We are very grateful to Dr. Laibale Friedman (Tel Aviv, Israel), Dr. Alexander Puchkov
(Kiev, Ukraine), Mr. Jiirgen Wiesner (Wolfsburg, Germany), Mr. Igor’ Ovsyannikov,
Mr. Alexander Sokolov, Mr. Pavel Udovichenko and Mr. Eugeny Shankhiza (all Mos-
cow, Russia) who kindly has given material and information for our study. Special
thanks to Prof. David L. Pearson (Arizona State University, USA) for revising the Eng-
lish text. For the first author the study received financial support from the Ministry of
Education and Science of the Russian Federation (Project No 6.632.2014/K).

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